B. Foggi, M. A. Signorini & G. Rossi

The genus Festuca in Italy

Introduction

Abstract

Foggi, B., Signorini , M. A. & Rossi, G.: The genus Festuca in Italy. ~ Bocconea 16(1): 55-64. 2003. ~ ISSN 1120-4060.

HackeI 's Monographia Festucarum Europaearum (1882) can be considered the starting point of modem systematic studies on this criticaI genus in Europe. After his work, almost one hun­dred years of infonnation was summed up by Markgraf-Dannenberg (1980) in her treatment of the genus in Flora Europaea. Here, the total number of species had increased from 28 (Hacke1 (882) to 170. However, this was due more to the raising to the rank of species of many infra­specific taxa, than to the recognition of new enti ti es as a result of taxonomic investigations. During these hundred years, the increase of knowledge on the systematic of this genus was very unequal in different European countries: in some of them like France, many in-depth investi­gations were carried out, while in other countries Iike Italy, studies started Iater and were much Iess exhausti ve. Our research team has been carrying out taxonomic investigations on the genus Festuca in Italy for several years, making use of modem systematic tooIs, among which typification and study of type material play an important role . In many instances, the results of our researches Ied to remarkable taxonomic rearrangements within criticaI groups. Some examples are illustrated, conceming the F violacea group, the endemie F robustifolia, and the relations between F gracilior and F inops.

The starting point ofmodem systematic studies on the difficult genus Festuca in Europe can be considered the Monographia Festucarum Europaearum (Hackel 1882). Here for the first time anatomical characters ofthe transverse section oftiller leafblades were taken into consideration, in addition to the traditional morphological characters. In Hackel 's sys­tematic conception, the genus includes a relatively small number of species (28), articu­lated in a large number of infraspecific taxa: subspecies, varieties and subvarieties.

After this work, almost one hundred years of information on the systematics of Festuca was summed up by Markgraf-Dannenberg, in her treatment of this genus in Flora Europaea (Markgraf-Dannenberg 1980). Here, the total number of species had increased enormously, from the 28 reported by Hackel, to 170. However, this was not only due to the recognition of new entities as a result of taxonomic investigations in criticai groups, but

56 Foggi & al.: The genus Festuca in Italy

also to the raising to the rank of species of many infraspecific taxa (Fig. l). If the total number of different taxonomic enti ti es is considered, instead of the number of species, the increase is considerably less: from 168 to 219. It must be emphasized, however, that in this peri od of time the increase of knowledge on the systematics of Festuca had been very unegual in different European countries. In some of them (for instance in France and in French-speaking countries, in Spain, and in the British Isles), many in-depth investigations had been carri ed out using modern systematic methods; while in other countries, like Italy, studies started later and were much less exhaustive.

Let us turn onr attention to the situation in ltaly (Fig. 2) . In this country, Hackel recog-

250 I

Fig. 1. Systematic knowledge on the genus Festuca in Europe

60

70

60

50

40

30

20

10

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IT l'L Y Ililcl<c l IT l'L Y rlkukgl. -Dunn . IT f'l Y P~",,,lIi

Fig. 2. Systematic knowledge on the genus Festuca in Italy.

Bocconea 16( I) - 2003 57

nised 68 different taxonomic entltles: 15 species with 65 infraspecific taxa. In Flora Europaea, only 63 different taxa are reported (55 species with 21 infraspecific taxa), that is fewer · than those repOlied by Hackel. I n the most recent Italian Flora, Pignatti's Flora d'Italia, the genus Festuca was treated by Pignatti himself in co-operation with Markgraf­Dannenberg (Pignatti & Markgraf-Dannenberg 1982). Here, 68 different taxonomic enti ­ties are listed , (61 species with 13 infraspecific taxa). Apart from the different conception ofrank, in the space ofjust one hundred years, from 1882 to 1982, the number of taxa is unchanged. Consequently, one could imagine that alI the taxonomic problems of Italian fescues had already been so lved by Hackel ; but this is not the case. ActualIy, as we already stated, during these one hundred years in Italy systematic investigations on this genus (and on the whole family Poaceae) had been very few: less than lO contributions conceming Festuca had been published in this periodo During that same time, about 150 papers appeared, for instance, in France, where scholars like Saint-Yves, Litardière, Auquier, Kerguélen, Plonka, and more recently Portai have been investigating the systematic ofthis genus since the publication of Hackel 's monograph. As a consequence, the number of dif­ferent taxa in France has been constantly increasing. In his recent treatment of the genus Festuca in France Portai reports for France 106 different taxa (Portai 1999), that is almost double the number reported by Hackel for the same territory 117 years before (Fig. 3). In ltaly, even if alI the contributions that have appeared since the publication of Pignatti's Flora are taken into consideration (Fig. 4), only the number of77 different taxa is reached. Yet, it is weII known that, generally speaking, the Italian flora is much richer than the French, thanks to a greater biogeographical diversity. Therefore, it sounds very strange that diversity within the genus Festuca is so much lower in Italy than in France. Even ifthis is possibly partly due also to the scarceness of floristic exploration of some parts of Italian territory, above ali it is related to the insufficient systematic investigation of this genus. After Hackel's monograph , few new taxa have been described for Italy, and most of them

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Fig. 3. Systematic knowledge on the genus Festuca in France.

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OTQIQI tlum lier ot different taxs

58 Foggi & al.: The genus Festuca in ltaly

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Fig. 4. Total number of different taxa of Festuca in Italy.

at the rank of species; this means that systematic relationships among taxa have not been sufficiently investigated. Consequently, there is stilI much work to be done.

Since 1996, our research team has been carrying out taxonomic investigations on the genus Festuca in ltaly, with special attention to the so-called "thin-Ieaved fescues" belong­ing to subgen. Festuca. Our approach has involved the use ofthe following essential inves­tigation tools:

- typification and study of type-material; - nomenclatural restatement; - morphometric analysis of macro- and micro-morphological features observed on spec-

imens colIected in the whole distribution area; - collection of morphological and ecological data trom living populations; - observation of features on living plants grown under controlled conditions; - karyological investigations.

It must be stressed that typification and study oftype-material must always be the start­ing point of ali investigations, as a high number of taxa have been validly described, and many names have been later misapplied, with the consequence of a great nomenclatural messo

The results of our investigations led in many instances to remarkable taxonomic rearrangements within criticai groups. The following are just a few examples.

The Festuca violacea aggregate in the Alps and the Apennines

This topic was discussed in Signorini & Foggi 1997; Foggi, Rossi & Signorini 1998, 1999.

Among the taxa of this group reported by different authors for the considered area, three

Bocconea 16( l) - 2003 59

species are confined only to the Eastern Alps: F narica, F picturata, F nitida. For these, our data essentially confirmed the taxonomic treatment proposed in a recent systematical review (Pils 1980) and later kept by other authors.

On the other hand, for those taxa growing also or exc1usively in the Western Alps and the Apennines, the results of our investigations led to a complete taxonomic revisiono For this area, in Hackel 's monograph two taxa are reported (Fig. 5a): F rubra varo nigricans growing in the whole Alpine chain; and F rubra subsp. vialacea subvar. typica for the Western Alps and the Apennines. In this latter taxan, he inc1uded al so F puccinellii, a

- = F J1lbro subvar. typico (incl. F pllccmellii)

-- = F J1lbro varo nigricans

a)

'. '

c)

•. - = F mocrothero

- = F violoceo

-- = F puccinellii

b)

• = F violoceo subsp. violoceo

• = F vi%cea subp. ito/ico

• = F violoceo subsp. puccinellii

* = F me/onopsis

Fig. 5. Distribution area in the Alps and the Apennines ofthe taxa belonging to the Festuca violacea

aggregate; a) data from Hackel (1882); b) data from Markgraf-Dannenberg (1980) and Pignatti & Markgraf-Dannenberg (1982) ; c) data from Foggi, Rossi & Signorini (1999).

60 Foggi & al.: The genus Festuca in Italy

species originally described for some localities in the Northem Apennines (Parlatore 1850). In Flora d'Italia (Pignatti & Markgraf-Dannenberg 1982) and fundamentally al so in Flora Europaea (Markgraf-Danneneberg 1980), the situation is radically changed (Fig. 5b). F violacea is raised to the rank of species and is restricted to the W-Alps, while pop­ulations from the Centrai and Southern Apennines are referred to F macrathera, a taxon that had been described by Hackel for Bosnia-Rercegovina (Rackel in Beck 1887). No species is reported for the Northem Apennines. In the Alps, F rubra var. nigricans is raised to the rank of species, under the name F puccinellii, but the locus classicus of this species (in the Northem Apennines) is surprisingly excluded from its distribution area.

The results of our investigations, which included multi variate analyses on morphomet­ric data, led to a completely different taxonomic outline. Two species were found to occur, well differentiated both karyologically and morphologically: F melanopsis - that is the new name for F rubra varo nigricans - in the Alps, and F violacea in the W-Alps and the Apennines, subdivided into three subspecies (Fig. Se). F melanopsis is a hexaploid (2n=42), while ali the three subspecies of F violacea proved to be diploid (2n= 14). The two species can be easi1y distinguished mainly on the basis of the appearance of leaf sheaths, which are smooth and straight in F melanopsis and corrugated, with somewhat entangled old fibers in the three subspecies of F violacea.

Populations of F violacea from the Northem Apennines have been referred to subsp. puccinellii. They are well differentiated from F rubra var. nigricans growing in the Alps, to which the name F puccinellii had been misapplied by Markgraf-Dannenberg.

Populations from the Centrai and Southern Apennines, included by Markgraf­Dannenberg in F. macrathera, we have referred to the new endemie subspecies italica. Our studies on the type material of F. macrathera showed, in fact, that this is a complete­Iy different taxon, which is possibly more closely related to F nitida than to F violacea.

Festuca robustifolia

This endemie taxon was originally described (Hackel 1882) as F ovina subv. robusta, with a distribution area confined to Monteferrato near Prato (Tuscany, ltaly). Markgraf­Dannenberg raised it to the rank of species under the new name F robustifolia. In Flora Europaea and in Flora d'Italia, the systematic conception of this taxon is much broader than the originai one held by Hackel and its distribution area includes a large part of Italy and a lot of different habitats (F ig. 6a). As a consequence, many records of F robustifòlia from ali parts of Italy and from very different ecological situations have been reported in floristic and vegetational works from then ono In a recent review on this species (Foggi & Signorini 2001), after studying the type material ofHackel's subvar. robusta and selecting a lectotype, we carri ed out a study on herbarium specimens, natural populations from the locus classicus, and living plants grown under controlled conditions. On the basis of our investigations, we were able to give a full description of morphological, karyological and ecological characters of F robustifolia. The species can be identified on the basis of:

- stems scabrid in the upper part, under the panicle; - sheaths glabrous; - leave blades scabrid;

Bocconea 16(1) - 2003 61

- transverse section ofleafblades regularly ovate, with sclerenchyma in a regularly thick­ened continuous ring.

We could detect that many ofthe records of F robustifolia from different ltalian regions were to be referred to other taxa, morphologically similar, but distinct from F robustifolia both systematically and ecologically. Among these:

- F faevigata Gaudin, with glabrous stem and broken sclerenchyma, widespread in open grasslands on limestone, over 1000 m, 2n=56;

- F billyi Kerguélen & Plonka, vicarious to F faevigata, on siliceous substrata, growing in Vaccinium heaths and Apennines grasslands at high altitudes, 2n=42;

- F stricta Host subsp. trachyphylla (Hack.) Patzke ex Pils, with pubescent sheaths and sclerenchyma in 3 more or less confluent strands, widespread in open arid grasslands and synanthropic, to 1000 m, 2n=42;

- F riccerii Foggi & Graz. Rossi, described for populations growing in the Northern Apennines (Appennino Tosco-Emiliano) in mountain grasslands over 1600 m, on sand­stone, 2n=28;

- F apuanica Markgr.-Dann., endemic to open grasslands and rocky sites in the Apuan Alps, 2n=70;

- F gamisansii Kerguélen subsp. aethaliae Signorini & Foggi, endemic to Mount Capanne on the Island ofElba, 2n=70.

In this narrow conception, F robusti[ofia is currently known with certainty only for ophi­olitic and ultramafic outcrops of southern Tuscany (Fig. 6b).

* = focus classicus

b)

Fig. 6. Distribution area of Festuca robustifolia; a) data from Markgraf-Dannenberg (1980) and Pignatti & Markgraf-Dannenberg (1982); b) data from Foggi & Signorini (2001).

62 Foggi & al. : The genus Festuca in Italy

Festuca inops - Festuca gracilior

This study is currently in progresso Festuca inops was described for Liguria, in North-Westem Italy (De Notaris 1844).

Rackel considered it as a subvariety of F ovina varo glauca, with a distribution area in the N-Apennines (Liguria, Tuscany).

Festuca gracilior was described by Rackel (1882) as a subvariety of F ovina varo duriuscula, with a distribution area from Spain (Pyrenees) to SE-France, and Italy (Tuscany). It was later raised to the rank of species by Markgraf-Dannenberg (1978). In Rackel's conception, the two taxa were fundamentally distinguished on the ground of glaucous color of leaves, a characteristic that actually can vary even within single popula­tions and is consequently ofno systematic value. In Flora Europaea and in Flora d'Italia, the two species appear to be almost indistinguishable on the basis of macro- and micro­morphological characters (Fig. 7). According to Flora Europaea, F gracilior grows in

F. inops F. gracilior

habit densely caespit ose densely caespit ose

stem 19-25(50) , smoot h or

2 0- 35 cm , glab rous scabridulous

0.4-0 .8 cm wide, ± (0.45) 0 .5-0 .8 cm wide,

leaves ± smooth , somewhat smooth , pruinose

pru in ose

veins 7 7

sclerenchyma complet e ring contin uous or s lig ht Iy

int errupt ed ring

ribs 5 3

o . ;~ o b

~ l

leaf section (from O Pignatti & O J:o Markgraf- ~o

Dannenberg) o

O

minutely ci li ate, closed in g laorous or puoerulem ,

sheaths closed in t he lower 1/ 4-the lower 1/ 4-1/2

1/ 2

panicle 3 .5-6 .5 cm , rat her dense 4-7 .5 cm , rather dense

spikelets 6-7 .7 mm , glaucous 6 .5- 7 .5 mm , green

upper glume 2 .7-4 .6 mm 3 .5-4 .5 mm

lemma 3 .9-5 mm 4-4.4(5 .5) mm

awn 0 -1 mm 0 .2-1 .5 mm

Fig. 7. Morphological characters of Festuca inops and Festuca gracilior (data from Markgraf-Dalmenberg 1980; Pignatti & Markgraf-Dannenberg 1982).

Bocconea 16( I) - 2003 63

France and Italy and is excluded from Spain; F inops is endemie to Italy (N-Apennines). In Italy and France, both species were found to be diploid, with 2n=14 (Kerguélen 1975 ; Bechi & Miceli 1995; our own data).

In the most recent papers on the gen. Festuca in Spain (e.g. de la Fuente & Ortunez 1998), a F gracilior is reported for the eastem part of the country, but with 2n=28. According to the most recent French contributions (Kerguélen & Plonka 1989; Portai 1999), in France this species surprisingly disappears just beyond the border with Spain, and is substituted by F occitanica (Litard.) Auquier & Kerguélen, a species morphologi­cally close to F gracilior with 2n=28.

We are currently investigating ltalian and French diploid populations referred both to F inops and to F gracilior. Inter- and intra-populational variation of micro- and macro-mor­phological characters in specimens coming from the whole area (including type material) are being analyzed. Intrapopulational variation is going to be investigated also by molec­ular analyses (RAPD).

Further investigations will also be necessary to clarify systematic relationships of these diploid taxa with the tetraploid taxa growing in SW France (reported under the name F occitanica) and in E Spain (reported under the name F gracilior, sensu De La Fuente & Ortunez, and incIuding also F tarraconensis (Litard.) Romo and F valentina (St.-Yves) Markgr.-Dann.).

Acknowledgements

We are grateful to Anne Maury for her kind revision of the English version of the papero

References

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64 Foggi & al. : The genus Festuca in Italy

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Ostalpenraum. - Pl. Syst. Evol. 136: 73- 124. Portai , R. 1999: Festuca de France. - Ed. Portai , Vals-près-Le-Puy. Signorini , M. A. & Foggi, B. 1997: A survey of the genus Festuca L. (Poaceae) in Italy. ilI.

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Addresses of the authors: B. Foggi , Museo di Storia Naturale sez. Orto Botanico, Università di Firenze, via P. A. Micheli 3, 1-5012 1 Firenze, ltaly. M. A. Signorin i, Dipartimento Biologia Vegetale, Università di Firenze, p.le delle Cascine 18, 1-50144 Firenze, Italy. G. Rossi, Dipartimento di Scienze del Territorio, Univers ità di Pavia, via S. Epifa ni o, 30,1-27100, Pav ia, Italy.